Radim Blažek

Rapid growth, early maturation and short generation time in African annual fishes.

BackgroundExtreme environmental conditions can give rise to extreme adaptations. We document growth, sexual maturation and fecundity in two species of African annual fish inhabiting temporary savanna pools.ResultsNothobranchius kadleci started to reproduce at the age of 17 days and size of 31 mm and Nothobranchius furzeri at 18 days and 32 mm. All four study populations demonstrated rapid growth rates of up to 2.72 mm/day (23.4% of their total length). Both species may produce diapausing embryos or embryos that are able to hatch in as few as 15 days, resulting in a minimum generation time as short as only one month. Incubation on the surface of damp peat moss results in high embryo survival (73%) and a high proportion of rapidly developing embryos (58%) that skip diapauses and hatch in less than 30 days. We further demonstrated that rapid growth and maturation do not compromise subsequent fecundity.ConclusionsOur data suggest that both species have the most rapid sexual maturation and minimum generation time of any vertebrate species, and that rapid maturity does not involve paedogenesis.

Parallel evolution of senescence in annual fishes in response to extrinsic mortality

BackgroundEarly evolutionary theories of aging predict that populations which experience low extrinsic mortality evolve a retarded onset of senescence. Experimental support for this theory in vertebrates is scarce, in part for the difficulty of quantifying extrinsic mortality and its condition- and density-dependent components that –when considered- can lead to predictions markedly different to those of the “classical” theories. Here, we study annual fish of the genus Nothobranchius whose maximum lifespan is dictated by the duration of the water bodies they inhabit. Different populations of annual fish do not experience different strengths of extrinsic mortality throughout their life span, but are subject to differential timing (and predictability) of a sudden habitat cessation. In this respect, our study allows testing how aging evolves in natural environments when populations vary in the prospect of survival, but condition-dependent survival has a limited effect. We use 10 Nothobranchius populations from seasonal pools that differ in their duration to test how this parameter affects longevity and aging in two independent clades of these annual fishes.ResultsWe found that replicated populations from a dry region showed markedly shorter captive lifespan than populations from a humid region. Shorter lifespan correlated with accelerated accumulation of lipofuscin (an established age marker) in both clades. Analysis of wild individuals confirmed that fish from drier habitats accumulate lipofuscin faster also under natural conditions. This indicates faster physiological deterioration in shorter-lived populations.ConclusionsOur data provide a strong quantitative example of how extrinsic mortality can shape evolution of senescence in a vertebrate clade. Nothobranchius is emerging as a genomic model species. The characterization of pairs of closely related species with different longevities should provide a powerful paradigm for the identification of genetic variations responsible for evolution of senescence in natural populations.

Alternative intrapopulation life-history strategies and their trade-offs in an African annual fish

In ephemeral habitats, the same genotypes cope with unpredictable environmental conditions, favouring the evolution of developmental plasticity and alternative life‐history strategies (ALHS). We tested the existence of intrapopulation ALHS in an annual killifish, Nothobranchius furzeri, inhabiting temporary pools. The pools are either primary (persisting throughout the whole rainy season) or secondary (refilled after desiccation of the initial pool), representing alternative niches. The unpredictable conditions led to the evolution of reproductive bet‐hedging with asynchronous embryonic development. We used a common garden experiment to test whether the duration of embryonic period is associated with post‐embryonic life‐history traits. Fish with rapid embryonic development (secondary pool strategy, high risk of desiccation) produced phenotypes with more rapid life‐history traits than fish with slow embryonic development (primary pool strategy). The fast fish were smaller at hatching but had larger yolk sac reserves. Their post‐hatching growth was more rapid, and they matured earlier. Further, fast fish grew to a smaller body size and died earlier than slow fish. No differences in fecundity, propensity to mate or physiological ageing were found, demonstrating a combination of plastic responses and constraints. Such developmentally related within‐population plasticity in life history is exceptional among vertebrates.

The interaction between an introduced fish host and local parasite fauna: Neogobius kessleri in the middle Danube River

Parasite communities of introduced fish Neogobius kessleri Günther (Gobiidae) were studied at five localities in the Slovak section of the Danube River during 2002–2005. Thirty-three metazoan parasite species were identified. All fish were infected with at least two parasite species; most of the parasite species were generalists. At all sampling sites, high susceptibility to local parasites was observed. The parasite community was dominated by three parasite species: glochidia of Anodonta anatina, larval or subadult acanthocephalan Pomphorhynchus laevis, and larval nematode Raphidascaris acus. The infection of both A. anatina and P. laevis was affected by season and habitat type, with higher abundance in spring and more frequent occurrence of A. anatina in side channels and P. laevis in main river channels. At both the component and infracommunity levels, a more diverse parasite community was found in side channels. This habitat was dominated by actively transmitted parasites, whilst endoparasites were more abundant in fish from the main river channel. Larval stages of parasites dominated the endoparasite community at all sampling sites. The introduced N. kessleri was used as intermediate host for most of the recorded parasites, in some cases also as a paratenic host. Finally, the importance of gobies as suitable hosts for local non-native parasite species (Anguillicoloides crassus, Anodonta woodina, Hydrozetes lacustris) is discussed.

Female bias in the adult sex ratio of African annual fishes: interspecific differences, seasonal trends and environmental predictors

Many populations have consistently biased adult sex ratios with important demographic and evolutionary consequences. However, geographical variation, the mechanisms, temporal dynamics and predictors of biased sex ratios are notoriously difficult to explain. We studied 334 wild populations of four species of African annual fish (Nothobranchius furzeri, N. kadleci, N. orthonotus, N. rachovii) across their ranges to compare their adult sex ratio, its seasonal dynamics, interpopulation variation and environmental predictors. Nothobranchius populations comprise a single age cohort and inhabit discrete isolated pools, with wide-ranging environmental conditions (habitat size, water turbidity, structural complexity, predators), making them ideal to study adult sex ratio variation. In captivity adult sex ratios were equal. In natural populations, adult sex ratios were biased 1:2 toward females in three study species while N. kadleci had sex ratios at unity. There was a decline in the proportion of males with age in one species, but not in the other species, implying most severe male mortality early after maturation, declining later perhaps with a decrease in male abundance. In general, the populations at vegetated sites had relatively more males than populations at sites with turbid water and little vegetation. Selective avian predation on brightly coloured male fish likely contributed to female dominance and vegetation cover may have protected males from birds. In addition, an aquatic predator, a large belastomid hemipteran, decreased the proportion of males in populations, possibly due to greater male activity rather than conspicuous colouration. Alternative explanations for a sex ratio bias, stemming from male–male contests for matings, are discussed. We conclude that the effect of environmental conditions on adult sex ratio varies dramatically even in closely related and ecologically similar sympatric species. Therefore, difficulties in explaining the ecological predictors of sex ratio biases are likely due to high stochasticity rather than limited sample size.

Laboratory breeding of the short-lived annual killifish Nothobranchius furzeri

Turquoise killifish, Nothobranchius furzeri, have an intrinsically short life span, with a median life span of <6 months and a maximum (90%) life span of 9 months. This short life span, which is unique among vertebrates, evolved naturally and has resulted in N. furzeri becoming a widely used laboratory model species in aging research and other disciplines. Here, we describe a protocol for the maintenance and breeding of the species under laboratory conditions. We provide details for egg incubation, hatching, everyday care of juvenile and adult fish, breeding and treatment of most common diseases. Emphasis is given to the fact that the requirements of N. furzeri substantially differ from those of other fish model taxa; N. furzeri live brief lives and in nature undergo nonaquatic embryo development, with consequences for their laboratory culture.

Trophic niche partitioning in communities of African annual fish: evidence from stable isotopes

Annual killifish of the genus Nothobranchius often co-occur in temporary savannah pools. Their space- and time-limited environment does not allow for any substantial habitat or temporal segregation. Coexisting species are therefore predicted to have well separated trophic niches to avoid intense food competition. Although in a previous “snapshot” study using stomach content analysis (SCA), the trophic niches of three sympatric species (N. furzeri, N. orthonotus, and N. pienaari) were found to vary among species, the difference was relatively weak and inconsistent across different sites. Here, we used the time-integrative capacity of stable isotope analysis to test whether the trophic niches of sympatric Mozambican Nothobranchius are more distinct over a long-term period. Analysis of carbon and nitrogen stable isotopes separated the trophic niche and trophic position of N. pienaari but failed to find any difference between N. furzeri/N. kadleci and N. orthonotus. No segregation was found at the sites with low prey diversity. In contrast, SCA identified N. orthonotus as the species with the most distinct trophic niche. We discuss the effect of prey diversity and different sensitivities of stomach content and stable isotope analysis in general and conclude that the trophic niches of the three sympatric Nothobranchius species are well separated.

Repeated intraspecific divergence in life span and aging of African annual fishes along an aridity gradient.

Life span and aging are substantially modified by natural selection. Across species, higher extrinsic (environmentally related) mortality (and hence shorter life expectancy) selects for the evolution of more rapid aging. However, among populations within species, high extrinsic mortality can lead to extended life span and slower aging as a consequence of condition‐dependent survival. Using within‐species contrasts of eight natural populations of Nothobranchius fishes in common garden experiments, we demonstrate that populations originating from dry regions (with short life expectancy) had shorter intrinsic life spans and a greater increase in mortality with age, more pronounced cellular and physiological deterioration (oxidative damage, tumor load), and a faster decline in fertility than populations from wetter regions. This parallel intraspecific divergence in life span and aging was not associated with divergence in early life history (rapid growth, maturation) or pace‐of‐life syndrome (high metabolic rates, active behavior). Variability across four study species suggests that a combination of different aging and life‐history traits conformed with or contradicted the predictions for each species. These findings demonstrate that variation in life span and functional decline among natural populations are linked, genetically underpinned, and can evolve relatively rapidly.

Population-specific responses to an invasive species

Predicting the impacts of non-native species remains a challenge. As populations of a species are genetically and phenotypically variable, the impact of non-native species on local taxa could crucially depend on population-specific traits and adaptations of both native and non-native species. Bitterling fishes are brood parasites of unionid mussels and unionid mussels produce larvae that parasitize fishes. We used common garden experiments to measure three key elements in the bitterling–mussel association among two populations of an invasive mussel (Anodonta woodiana) and four populations of European bitterling (Rhodeus amarus). The impact of the invasive mussel varied between geographically distinct R. amarus lineages and between local populations within lineages. The capacity of parasitic larvae of the invasive mussel to exploit R. amarus was higher in a Danubian than in a Baltic R. amarus lineage and in allopatric than in sympatric R. amarus populations. Maladaptive oviposition by R. amarus into A. woodiana varied among populations, with significant population-specific consequences for R. amarus recruitment. We suggest that variation in coevolutionary states may predispose different populations to divergent responses. Given that coevolutionary relationships are ubiquitous, population-specific attributes of invasive and native populations may play a critical role in the outcome of invasion. We argue for a shift from a species-centred to population-centred perspective of the impacts of invasions.

An overview of the Gyrodactylus (Monogenea: Gyrodactylidae) species parasitizing African catfishes, and their morphological and molecular diversity.

An overview of Gyrodactylus infecting catfishes from the African continent is provided, including new data from Sudan, Senegal, Kenya and Mozambique. Haptoral sclerite morphometry and nuclear ribosomal DNA sequences revealed the presence of eight Gyrodactylus species. On Senegalese Synodontis nigrita, Gyrodactylus synodonti n. sp. and Gyrodactylus nigritae n. sp. are described. These are the first reports of gyrodactylid parasites from mochokid hosts. From the fins of North African catfish Clarias gariepinus collected in Mozambique, Gyrodactylus alekosi n. sp. and Gyrodactylus rysavyi were identified. G. rysavyi was also reported from Kenyan C. gariepinus and Senegalese Clarias anguillaris. From the fins of C. anguillaris studied in Senegal, two more species, Gyrodactylus transvaalensis and Gyrodactylus gelnari n. sp. were recognised. In addition, Gyrodactylus turkanaensis n. sp. from the gills of Kenyan C. gariepinus was described and an undescribed Gyrodactylus sp. was recorded from Sudanese representatives of the same host. Detailed morphometrical and molecular comparisons of the species are presented and discussed. The study highlights the hitherto understudied diversity of viviparous monogenean parasites throughout Africa.